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Salp

Salps are barrel-shaped, gelatinous planktonic tunicates in the family Salpidae, subphylum Tunicata, and phylum Chordata, making them distant relatives of vertebrates such as humans and fish rather than jellyfish.^[1]^[2] These free-floating marine invertebrates, typically ranging from a few millimeters to over 20 centimeters in length, inhabit open ocean waters globally and are known for their distinctive life cycle alternating between solitary individuals and colonial chains.^[3]^[4] Salps propel themselves through the water column using a jet-like mechanism, contracting their transparent, muscular bodies to pump water in through an opening at one end and expel it from the other, achieving speeds of up to 30 body lengths per minute while filtering vast volumes of seawater.^[4]^[5] As efficient filter feeders, they strain phytoplankton, bacteria, and other microscopic particles through internal mucous nets, filtering over 1,000 times their body volume per hour and playing a pivotal role in marine food webs by transferring energy from primary producers to higher trophic levels.^[6]^[7]^[8] Their complex anatomy includes a dorsal nerve cord and functional organs like a heart, pharynx, and endostyle, underscoring their evolutionary significance as basal chordates with more advanced systems than many other gelatinous zooplankton.^[2] Ecologically, salps are key contributors to the ocean's biological carbon pump, forming dense blooms that rapidly sink fecal pellets and mucous houses to the deep sea, potentially sequestering substantial amounts of atmospheric carbon dioxide and mitigating global warming effects.^[9]^[10] These blooms can dominate pelagic ecosystems during certain seasons, influencing nutrient cycling, microbial communities, and even fisheries by outcompeting other grazers like krill.^[11]^[12]

Biology

Morphology and Anatomy

Salps possess a distinctive barrel-shaped, gelatinous body enclosed in a transparent test composed primarily of cellulose, which provides structural support and buoyancy while remaining flexible.^[13] Typical body lengths range from 1 to 10 cm, although some species, such as those in the genus Salpa, can attain lengths up to 30 cm.^[14] The test is secreted by the underlying epithelium and consists of an outer layer of protein and an inner tunic of cellulose microfibrils embedded in a mucoid matrix, rendering the animal nearly transparent and aiding in camouflage within the water column.^[13] The body features two prominent siphons at opposite ends: an anterior oral siphon for water intake and a posterior atrial siphon for expulsion.^[15] Encircling the cylindrical body are 6 to 10 circular muscle bands that enable rhythmic contractions, drawing water into the body cavity through the oral siphon for feeding and respiration, then expelling it forcefully through the atrial siphon.^[15] This muscular arrangement supports pulsatile jet propulsion, allowing salps to achieve mean swimming speeds of 1.2-1.7 cm/s (equivalent to ~0.3 body lengths/s), with jet pulses up to ~3.3 cm/s, efficiency enhanced by the streamlined barrel shape that minimizes drag.^[16] Internally, salps exhibit a simplified anatomy typical of tunicates, including a central gut that forms a compact loop extending from the pharynx to the anus near the atrial siphon.^[17] The pharyngeal region houses a ventral endostyle, a glandular structure that secretes mucus to trap phytoplankton and other particles, and an expansive branchial basket lined with numerous gill slits (stigmata) that facilitate filter-feeding by straining food from incoming water currents.^[18] Filtered water and waste are then directed into the surrounding atrial cavity before ejection via the posterior siphon, completing both the feeding and propulsive cycles.^[13] The nervous system is simple, consisting of a cerebral ganglion connected to peripheral nerves that coordinates muscle contractions and sensory responses.^[17] Sensory structures include simple ocelli positioned dorsally for phototaxis, enabling light detection to guide vertical migration and predator avoidance.^[17] Morphological variations occur between the solitary (oozooid) and aggregate (blastozooid) life stages, with solitary forms generally larger and more robust to support sexual reproduction, while aggregate forms are smaller and form chains for asexual budding.^[19] These differences in size and arrangement influence locomotion, as solitary salps rely on individual jet pulses, whereas aggregates synchronize contractions for coordinated chain movement.^[20]

Life Cycle and Reproduction

Salps possess a biphasic life cycle featuring obligatory alternation between solitary (oozooid) and colonial (blastozooid) generations, enabling both rapid population expansion and genetic recombination. The solitary oozooid stage initiates the cycle by reproducing asexually through stolon budding, where a long stolon extends from the posterior of the parent and develops into a chain of genetically identical blastozooids that are released as an aggregate colony. This asexual phase allows for exponential proliferation, as a single oozooid can produce dozens to hundreds of blastozooids in a single chain.^[21]^[22] In the aggregate blastozooid stage, reproduction is sexual and hermaphroditic; younger (female-phase) individuals in the chain are cross-fertilized by sperm from older (male-phase) individuals in the same or nearby chains, with fertilization typically occurring in the atrial cavity. A single embryo develops within the ovary of a blastozooid, nurtured via a placental connection until it matures into a new oozooid, which is then released from the chain. Aggregate chains vary by species but can comprise hundreds of individuals and extend up to several meters in length, facilitating collective swimming and feeding efficiency.^[23]^[22]^[24] The developmental timeline from embryo to mature solitary oozooid typically spans several weeks, with growth rates strongly influenced by environmental factors such as temperature and food availability; warmer waters and abundant phytoplankton accelerate maturation, supporting swift generational turnover. Asexual reproduction in the oozooid stage confers advantages for rapid colonization of nutrient-rich patches, potentially leading to population booms, whereas sexual reproduction in the blastozooid stage introduces genetic diversity to enhance adaptability to varying conditions.^[24]^[25]^[21] Mortality in salps is stage-specific: aggregate chains face heightened predation risk from fish and other marine predators due to their conspicuous linear formations, while solitary oozooids are more vulnerable to starvation during periods of low phytoplankton abundance, as their larger size demands sustained filter-feeding. These factors contribute to the cyclical nature of salp populations, with high reproductive output offsetting losses.^[25]^[26]

Taxonomy and Evolution

Classification

Salps are classified within the phylum Chordata, subphylum Tunicata, class Thaliacea, and order Salpida, encompassing two families: Salpidae, which includes 9 genera and approximately 70 species, and Cyclosalpidae, with 1 genus and about 10 species.^[27]^[28] This hierarchical placement reflects their position as pelagic tunicates closely related to other chordates, distinguished from sessile ascidians by their free-swimming lifestyle.^[29] Key genera within these families illustrate the diversity of salp forms and distributions; for instance, the genus Salpa in the Salpidae family is cosmopolitan and includes species like Salpa maxima, known for extensive chain formations in temperate and subtropical waters, while Cyclosalpa in the Cyclosalpidae family inhabits deeper waters, and Ihlea is prominent in polar regions such as the Southern Ocean.^[27]^[30] Species within Salpida are primarily distinguished by morphological traits such as the arrangement of individuals in linear or circular chains, the structure of the gelatinous test (outer tunic), and variations in siphon morphology, which facilitate jet propulsion; in total, around 80 species have been described, with molecular methods continuing to reveal additional diversity.^[31] The taxonomic framework for salps originated with Jean-Baptiste Lamarck's 1816 classification of tunicates, where he first grouped salps separately from mollusks based on their anatomical features. Subsequent refinements in the 20th century, particularly through the application of electron microscopy by researchers like Rob W.M. van Soest, enabled detailed examination of internal structures and led to revisions in generic boundaries and species delineations. Currently, taxonomic challenges persist due to cryptic species complexes, which have been increasingly identified through DNA barcoding techniques since the 2010s, highlighting intraspecific genetic variation that morphological criteria alone cannot resolve.^[31]

Phylogenetic Relationships

Salps belong to the subphylum Tunicata within the phylum Chordata, positioning them as invertebrate chordates and the closest living relatives to vertebrates. Like all tunicates, salps exhibit key chordate synapomorphies during their larval stage, including a notochord for structural support and a dorsal hollow nerve cord that coordinates basic sensory and motor functions, features shared with vertebrates but lost or modified in the adult form. Within Tunicata, salps are part of the class Thaliacea, which is derived alongside the sessile Ascidiacea (sea squirts) and planktonic Appendicularia (larvaceans), forming a monophyletic group that diverged from other chordates early in evolutionary history.^[32]^[33] Molecular phylogenetic studies using 18S rRNA and mitochondrial DNA sequences from the 2000s onward have clarified the position of Thaliacea as a monophyletic clade and sister group to Appendicularia within Tunicata. Analyses of 18S rDNA sequences indicate that Thaliacea diverged from Appendicularia approximately 450 million years ago during the Ordovician-Silurian transition, with molecular clock estimates around 447 million years ago (95% CI: 411–484 Mya); internal relationships among thaliacean orders (Salpida, Doliolida, and Pyrosomida) showing Pyrosomida as the earliest-branching lineage. These findings, supported by mitochondrial cytochrome oxidase I (cox1) data, resolve Thaliacea as a cohesive unit distinct from Ascidiacea, though early studies highlighted challenges due to the rapid evolutionary rate of thaliacean 18S rRNA sequences.^[30]^[34]^[35]^[32] The fossil record of tunicates, including salps, is sparse due to their soft-bodied nature, with salp-specific preservation particularly rare and mostly inferred from modern analogs rather than direct fossils. The earliest tunicate-like fossils date to the mid-Cambrian around 500 million years ago, such as the soft-tissue preserved Megasiphon thylakos, which suggests an ascidiacean-like body plan with filter-feeding adaptations. Later Ordovician evidence (~450 million years ago) includes bioimmured traces in bryozoans interpreted as tunicate holdfasts, supporting a deep origin for Tunicata but providing no unambiguous salp remains.^[36] In broader animal relationships, salps display a simplified nervous system compared to vertebrates—lacking a brain and relying on a diffuse nerve net—yet feature advanced jet-propulsion locomotion via muscular contractions, reflecting pelagic adaptations within chordates. Debates persist on the monophyly of Tunicata, with phenotypic and molecular data overwhelmingly supporting it as a clade sister to Vertebrata, though some early morphological analyses suggested paraphyly. Recent post-2020 genomic studies have revealed fragmented Hox gene clusters in tunicates, including salps, that parallel vertebrate clusters in gene content but lack colinearity, reinforcing their shared chordate ancestry while highlighting extensive genomic rearrangements early in the lineage.^[37]^[33]^[38]

Distribution and Habitat

Global Range

Salps exhibit a cosmopolitan distribution across all major ocean basins, from polar to tropical waters, inhabiting epipelagic zones typically between 0 and 200 meters depth, though they are notably absent from brackish or freshwater environments.^[26]^[39]^[40] Their highest species diversity occurs in temperate and subtropical regions, where warmer waters support a greater variety of salp genera and life stages compared to polar or highly oligotrophic areas.^[41]^[42] In the Northern Hemisphere, salps form notable concentrations in the North Atlantic, particularly through periodic blooms of Salpa aspera in the Slope Water region south of New England, where swarms can cover extensive areas during summer months.^[43] In the Southern Ocean, swarms are prominent, with species such as Ihlea racovitzai distributed widely north of approximately 64°S in regions like the Lazarev Sea, and Salpa thompsoni dominating Antarctic epipelagic communities.^[44] Indo-Pacific waters host significant concentrations, including diverse assemblages in the central Pacific, where temperature gradients influence species like Salpa fusiformis across subtropical latitudes.^[40]^[39] As of late 2024, unprecedented salp events were observed along southeastern Tasmanian beaches and bays, highlighting variability in temperate distributions.^[45] Salps commonly undertake diurnal vertical migrations, descending to depths of up to 800 meters during the day to evade visual predators and ascending toward the surface at night to access phytoplankton-rich layers, with migration amplitudes varying by species and region.^[46]^[47]^[48] For instance, Salpa thompsoni in the Southern Ocean migrates from daytime depths around 450 meters to nighttime positions near 100 meters, enhancing foraging efficiency while minimizing predation risk.^[49] While most salp species are widely distributed with low endemism, a few act as polar specialists, such as Salpa thompsoni, which is predominantly confined to Antarctic waters and plays a key role in high-latitude ecosystems.^[50]^[51] Ongoing monitoring through satellite-derived chlorophyll anomalies and targeted net tows has revealed range expansions for several salp species since the early 2000s, with increased detections in subpolar and transitional zones via large-scale surveys like the CCAMLR 2000 expedition in the Atlantic sector of the Southern Ocean.^[52]^[53]^[54]

Environmental Adaptations

Salps demonstrate broad physiological tolerance to key marine environmental variables, enabling their widespread distribution across oceans. They are predominantly eurythermal, with species inhabiting waters ranging from near-freezing polar temperatures around 0°C to subtropical conditions up to approximately 30°C. For instance, in the temperate species Salpa fusiformis, routine metabolic rates increase markedly with temperature, from 1.66 μmol O₂ g⁻¹ h⁻¹ at 10°C to 3.95 μmol O₂ g⁻¹ h⁻¹ at 17°C, yielding a Q₁₀ value of 3.45 that reflects heightened physiological activity in warmer conditions.^[55] Salps are adapted to typical oceanic salinities of 30–40 ppt, though specific tolerance limits remain understudied; deviations beyond this range can stress osmoregulatory processes in these gelatinous organisms. Additionally, they exhibit notable hypoxia tolerance, consuming oxygen down to undetectable levels in experimental conditions at both 10°C and 17°C before recovering upon reoxygenation, which facilitates diel vertical migrations into oxygen-minimum zones such as those in the California Current.^[55]^[56] Feeding adaptations further enhance salps' resilience in fluctuating nutrient environments. As passive filter feeders, they efficiently process phytoplankton-rich water through mucous nets, with grazing rates removing up to 24.5% of daily primary production in bloom conditions and exhibiting peak efficiency when particle concentrations align with optimal mesh sizes (typically 1–10 μm). Gut passage times are rapid, averaging 1–2 hours depending on body size and food density, as calculated from pigment clearance and marking experiments, allowing swift nutrient assimilation and fecal pellet formation even at low food levels. In nutrient-poor waters, salps adapt by downregulating metabolism and reducing activity, conserving energy stores derived from prior blooms.^[57]^[58]^[59] Buoyancy regulation is achieved primarily through their low-density, gelatinous composition, with body water content exceeding 95%, conferring near-neutral buoyancy that minimizes energy expenditure for suspension in the water column. Unlike some plankton, salps lack gas vacuoles or prominent lipid inclusions for buoyancy; instead, their transparent tunic and internal fluid dynamics provide passive stability. In colonial forms, chain orientation aligns with currents via hydrodynamic forces, optimizing position relative to food patches or avoiding sinking.^[60] Sensory capabilities support behavioral responses to environmental gradients. Phototaxis is mediated by simple ocelli and dispersed photoreceptors that hyperpolarize in response to light, guiding vertical migrations toward surface phytoplankton during daylight and deeper waters at night. Geotaxis, detected via statocysts or body orientation cues, aids in maintaining upright posture and chain alignment against gravity. Bioluminescence is absent in salp taxa, distinguishing them from bioluminescent relatives like pyrosomes; however, structural iridescence in some species, arising from tunic scattering, contributes to optical camouflage by blending with ambient light fields and reducing visibility to predators.^[61]^[62] Under stress from low-food conditions, salps adapt behaviorally by extending chain lengths, sometimes reaching several meters, to amplify collective filtration volume and enhance encounter rates with sparse particles—a response that boosts foraging success without increasing individual energy costs. This plasticity, combined with siphon-mediated jet propulsion for fine adjustments, underscores their efficiency in oligotrophic open-ocean settings.^[63]

Ecology and Interactions

Role in Food Webs

Salps function as primary consumers in marine food webs, primarily grazing on phytoplankton, bacteria, and detritus through their filter-feeding mechanism.^[64] This feeding strategy allows them to process large volumes of seawater, with large solitary individuals capable of clearing up to tens of liters per day, and clearance rates exceeding 100 liters per individual per day reported in some species during high-activity periods.^[65]^[66] By consuming these microbial and particulate resources, salps help regulate microbial loop dynamics and contribute to the transfer of low-level primary production upward in the trophic structure.^[63] As prey, salps occupy a key position for higher trophic levels, serving as food for various fish species such as herring and mackerel, seabirds, certain baleen whales that opportunistically target dense swarms, and some invertebrates including medusae.^[67]^[3]^[51] However, their high water content, approximately 95%, results in low nutritional value, making them a less energetically rewarding food source compared to crustacean prey like krill.^[51] This gelatinous composition limits their role as a sustained energy provider for predators requiring high-calorie diets.^[26] Salps enhance trophic efficiency by converting dilute phytoplankton energy into biomass accessible to predators, potentially increasing overall energy transfer in ecosystems during their abundance.^[68] Yet, their short life spans, often lasting only weeks to months, constrain long-term biomass accumulation and efficient energy passage to higher levels.^[69] Additionally, salps engage in occasional symbiotic interactions, such as associations with hyperiid amphipods that may provide mutual benefits including nutrient supplementation, and harbor gut bacteria that aid in processing low-nutrient diets.^[70]^[71] In terms of competition, salps vie with copepods and krill for phytoplankton resources, potentially displacing these crustaceans during bloom events through superior filtration capacities.^[72]

Bloom Dynamics and Predation

Salp blooms are primarily triggered by environmental conditions that enhance phytoplankton availability, such as nutrient upwelling from coastal or deep waters, which fuels the initial food source for salps. This abundance promotes the asexual blastozooid phase of their life cycle, where embryos develop into chains of genetically identical individuals, enabling exponential population growth through repeated budding. Warm water temperatures, often associated with seasonal warming or oceanographic anomalies, further accelerate this reproductive mode by optimizing metabolic rates and reducing developmental times. During peak blooms, salp densities can surpass 1000 individuals per cubic meter, transforming sparse populations into dense swarms within days.^[73]^[74]^[75] The spatial and temporal dynamics of salp blooms vary by region but typically encompass vast areas and extended periods. Swarms can extend over thousands of square kilometers, with documented historical events reaching up to 100,000 km² in extent. For instance, a bloom of Thalia democratica in the California Current covered approximately 9,000 km². These events often persist for weeks to months, driven by the interplay of advection by currents and sustained favorable conditions, before declining due to food exhaustion or aggregation into sexual oozooids that initiate the next cycle. Global hotspots include upwelling zones like the California Current and eastern Australia, where such dynamics recur seasonally.^[76]^[77]^[74] Predation on salps involves both specialized and opportunistic marine predators, influencing bloom persistence. Specialized fish like the American harvestfish (Peprilus paru) target salps as a primary food source, using their streamlined bodies to pursue gelatinous prey efficiently. Generalist predators, such as the ocean sunfish (Mola mola), also consume salps opportunistically during blooms. In response, salps exhibit anti-predator adaptations, including rapid jet propulsion for escape speeds up to 20 body lengths per second and synchronized contractions within chains to achieve steadier, faster group locomotion. Chain fragmentation may occur under duress, allowing individuals to disperse and evade capture.^[78]^[79]^[80]^[81] Blooms exert significant short-term ecological effects through intense grazing pressure. High salp densities rapidly deplete surface phytoplankton biomass, often reducing chlorophyll a concentrations by over 60% in affected areas and shifting energy flow away from crustacean grazers. This depletion disrupts microbial loops by limiting dissolved organic matter available to bacteria, potentially favoring smaller heterotrophs adapted to low-nutrient conditions. Post-bloom, the mass sinking of salp carcasses enhances passive carbon export to mesopelagic depths, sequestering organic material from surface waters.^[52]^[82] Modeling salp bloom dynamics relies on structured population approaches to simulate growth trajectories without exhaustive parameterization. Age- or stage-based matrix models, akin to Leslie frameworks, incorporate asexual reproduction rates to qualitatively depict exponential phases driven by chain elongation, peaking at high densities before resource-driven collapse. These models highlight sensitivity to initial aggregate presence and environmental cues, aiding predictions of bloom onset in prone regions like subtropical fronts. Individual-based simulations further refine these patterns by accounting for spatial dispersion and mortality.^[83]^[84]

Ecological and Oceanographic Importance

Carbon Cycling and Nutrient Dynamics

Salps play a significant role in the ocean's biological carbon pump through the production of dense fecal pellets and the sinking of dead chains, which facilitate the export of organic carbon from surface waters to the deep sea. These fecal pellets, formed from the efficient filter-feeding mechanism of salps, sink at rapid rates of 400–1,200 meters per day, allowing them to bypass much of the upper ocean remineralization zone.^[10] In areas of salp blooms, this process exports 10–25% of primary production to depths below 200 meters on average, with peaks reaching up to 46% during intense blooms, as measured by neutrally buoyant sediment traps deployed in the Southern Ocean.^[58] The sinking of entire dead salp chains further contributes to this flux, though fecal pellets account for the majority (up to 78%) of salp-mediated carbon export.^[85] In the Southern Ocean, salps are particularly influential, contributing 15–50% of the total carbon flux to intermediate depths in regions where they dominate over krill, based on fecal pellet carbon production estimates from moored sediment traps.^[86] In contrast to krill-dominated systems, where fecal pellets exhibit higher export efficiency (~72% to 300 m) but lower production, salp pellets contribute equally to flux at 300 meters despite producing four times more carbon than krill pellets, due to their greater abundance during blooms.^[86] Measurement techniques such as sediment traps and stable isotope tracing (e.g., δ¹³C) confirm that salp fecal pellets sink faster and retain more carbon than those from copepods, with salp pellets showing minimal isotopic fractionation indicative of rapid transit.^[87] Salps also influence nutrient dynamics through excretion and sloppy feeding, releasing ammonium and phosphate that stimulate bacterial growth in surface waters. Ammonia excretion rates in species like Salpa fusiformis increase allometrically with body size and exponentially with temperature, providing readily available nitrogen for microbial communities.^[88] Phosphate release occurs similarly via metabolic processes and incomplete particle capture during filter feeding, enhancing nutrient regeneration and potentially altering N:P ratios in salp-dominated areas compared to crustacean systems.^[89] However, salp fecal pellets remineralize slowly due to low microbial respiration rates (<1% of pellet carbon per day), preserving nutrients for deeper export rather than rapid recycling.^[10] Compared to other zooplankton, salps are more efficient carbon exporters owing to their large body size, which produces compact pellets, and low individual respiration rates that minimize carbon loss during grazing. This efficiency can increase the biological pump's transfer of net primary production to sinking particulate organic carbon by 1.5-fold in salp-influenced waters.^[10]

Impacts of Climate Change

Climate change is driving notable range shifts in salp populations, with observed poleward expansions since the 1990s, particularly in the Southern Ocean where warming oceans of 1-2°C have facilitated increased incursions into Antarctic waters previously dominated by other zooplankton. For instance, species like Salpa thompsoni have shown southward distribution expansions linked to rising sea surface temperatures and reduced sea ice cover, allowing salps to intrude into higher-latitude regions around the Antarctic Peninsula. Models project that these shifts could lead to a 20-50% increase in salp bloom frequency by 2100 under high-emission scenarios, potentially altering seasonal dynamics in polar and subpolar ecosystems.^[58]^[72]^[90] Physiologically, elevated temperatures accelerate salp metabolism and reproduction rates, enabling faster population growth in warming waters, though extreme heat can reduce individual survival and chain integrity. Studies indicate that metabolic rates in salps decrease with cooling but rise significantly under moderate warming, supporting higher energy demands for filter-feeding and locomotion.^[91]^[92] These changes are precipitating ecosystem shifts, notably in the Southern Ocean where salps are increasingly replacing Antarctic krill (Euphausia superba), disrupting food webs for higher trophic levels such as penguins, seals, and baleen whales that rely on krill. Enhanced salp abundance promotes greater carbon export to deep waters through fecal pellets and mucous nets, potentially amplifying the ocean's biological pump, but this transition risks fishery disruptions by reducing nutrient recycling and altering primary production efficiency.^[93]^[94]^[10] Despite these insights, salp responses to climate change remain understudied in tropical upwelling zones, where warming could interact with nutrient pulses to drive unpredictable blooms. Experts advocate for expanded monitoring using satellite remote sensing, bio-ARGO floats, and citizen science initiatives to track these dynamics and inform mitigation strategies, as current data gaps hinder accurate projections of ecosystem resilience.^[95]^[3]

History and Research

Discovery and Early Studies

Salps have been observed by seafarers for centuries, often referred to as "sea grapes" due to their gelatinous, grape-like appearance in chains floating on the ocean surface.^[96] Early scientific descriptions emerged in the mid-18th century, with Edward Browne providing one of the first accounts in 1756, followed by Peter Forsskål's observations in 1775 of their tubular, transparent forms in Mediterranean waters.^[26] The genus Salpa was formally established by Peter Forsskål in 1775, classifying them within the tunicates based on limited morphological data available at the time.^[97] Detailed anatomical studies advanced in the early 19th century, with Georges Cuvier offering a comprehensive description of salp structure in his 1804 Tableau élémentaire de l'histoire naturelle des animaux, distinguishing their barrel-shaped bodies and jet propulsion from other gelatinous organisms. Initially, salps were frequently misclassified as medusae (jellyfish) owing to their similar translucent, gelatinous forms, a confusion resolved only with improved microscopy revealing their chordate affinities and internal test structure by the mid-1800s.^[26] Regional investigations contributed to early knowledge, particularly in the Mediterranean where Italian naturalists documented salp occurrences during the 1700s, noting their seasonal abundances in coastal waters.^[26] Pacific explorations during James Cook's voyages in the 1770s also recorded gelatinous plankton resembling salps, though without formal identification, highlighting their widespread presence in open oceans.^[98] A pivotal milestone was the recognition of salps' alternation of generations, first proposed by Adelbert von Chamisso in 1819 and elaborated by Édouard van Beneden in the 1880s through dissections showing the shift between solitary asexual and colonial sexual phases.^[26] Concurrently, Victor Hensen's 1889 Plankton-Expedition linked salps to broader oceanographic patterns, using quantitative nets to sample them as key planktonic components during the first systematic Atlantic survey. The HMS Challenger expedition (1872–1876) marked a turning point by collecting the first global dataset of salp specimens across multiple oceans, as detailed in William Abbott Herdman's 1888 report on Tunicata, which cataloged species distributions and morphologies from over 300 stations. These findings spurred early 20th-century monographs, including William E. Ritter's studies on Pacific salp life cycles around 1905, emphasizing their reproductive strategies and ecological roles based on Scripps Institution collections.^[99]

Modern Research Advances

Modern research on salps has leveraged advanced technological tools to enhance detection and understanding of their biology and ecology. Since the 1990s, acoustic imaging techniques, such as multifrequency echosounders, have been employed to detect and quantify salp blooms by measuring backscattering from their gelatinous bodies, allowing non-invasive mapping of distributions in the water column.^[100] Similarly, genomic analyses have provided insights into salp evolutionary biology; a preliminary genome assembly for Salpa thompsoni in 2016 revealed rapid evolutionary rates and unique signatures compared to other chordates, while a full genome and transcriptome study in 2023 highlighted molecular adaptations, including abundant repetitive elements and G-quadruplex motifs, that support reproductive flexibility in changing environments.^[101]^[24] Key long-term studies have quantified salps' contributions to ocean processes. The Atlantic Meridional Transect (AMT) program, ongoing since 1995, has monitored salp abundances across latitudinal gradients, revealing their significant role in carbon export through fecal pellet production, with blooms potentially contributing substantially to particle flux in oligotrophic regions.^[102] In the 2020s, remote sensing technologies, including satellites and drones, have been integrated to track environmental drivers of blooms; for instance, post-El Niño conditions in 2023–2024 correlated with elevated salp densities off Southern California, detected via changes in sea surface conditions and chlorophyll anomalies.^[103] Interdisciplinary research links salp dynamics to broader systems. Climate models incorporating salp grazing have shown their influence on carbon sequestration, with blooms enhancing export efficiency in Southern Ocean simulations.^[10] Salp outbreaks also impact fisheries; in the 2010s, massive blooms of Ihlea magellanica in Chile's Chiloé Island region clogged salmon farm nets and led to fish mortality from overfeeding on salps.^[104] Emerging studies explore salp-associated microbial communities and pollutants. Microbiome research indicates that salps host specific bacterial symbionts, such as Psychrobacter species, which may facilitate cellulose digestion in their guts, enhancing nutrient processing during blooms.^[105] Investigations into plastic pollution have documented microplastic ingestion; every examined salp in North Pacific samples contained mini-microplastics (<333 μm) in their guts, reflecting ambient seawater concentrations and highlighting salps as bioindicators.^[106] Recent advances as of 2025 include detailed observations of salp colony swimming patterns, revealing coordinated jet propulsion that produces helical trajectories, as documented using advanced underwater imaging in 2024. Additionally, a 2025 study emphasized the outsized role of salps alongside other zooplankton in Southern Ocean carbon export, underscoring their contribution to global carbon cycling.^[107]^[108] Future research directions include predictive modeling and habitat assessments. Artificial intelligence approaches are being developed to forecast plankton blooms, including salps, by integrating satellite data and environmental variables for early warning systems.^[109] Additionally, evaluations of indirect threats from climate-driven habitat alterations, such as ocean warming and acidification, are assessing potential shifts in salp distributions and bloom frequencies.^[24]

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Salps are barrel-shaped marine invertebrates that swim by jet propulsion. Morphological variations among species and life-cycle stages are accompanied by ...
[20]
A New Molecular Phylogeny of Salps (Tunicata: Thalicea: Salpida ...
Salps are marine pelagic tunicates with a complex life cycle, including a solitary and colonial stage composed of asexually budded individuals.
[21]
AboutSalps - Florida State University
Salps have two different life stages: a solitary asexual stage and a colonial sexual stage. The solitary stage of salps (also referred to as the oozooid stage, ...Missing: reproduction biphasic
[22]
Reproduction and population structure of the salp Iasis zonaria ...
Apr 25, 2013 · The complex life history of salps involves an alternation between the solitary (oozooids) and aggregate (blastozooids) generations. Oozooids ...Missing: biphasic | Show results with:biphasic
[23]
Salpa genome and developmental transcriptome analyses reveal ...
Nov 29, 2023 · Our findings suggest salp are poised for reproductive success at birth, increasing the potential for bloom formation as ocean temperatures rise.Missing: biphasic | Show results with:biphasic
[24]
A review of the life cycles and life-history adaptations of pelagic ...
Oct 3, 2011 · The salp life cycle consists of two life-history stages and obligatory alternation of sexually and asexually reproducing generations ( ...Phylogenetic Relationships · Salps · DoliolidsMissing: biphasic | Show results with:biphasic<|control11|><|separator|>
[25]
An assessment of environmental and ecological drivers of salp ...
Dec 10, 2024 · They have a complex life history, with alternating generations between two reproductive stages: an asexual 'budding' solitary (oozoid) stage ...Missing: biphasic | Show results with:biphasic
[26]
WoRMS - World Register of Marine Species - Salpidae Lahille, 1888
### Taxonomic Hierarchy and Summary for Salpidae
[27]
https://www.marinespecies.org/aphia.php?p=taxdetails&id=137217
[28]
Report for eight species of Salpinae (Thaliacea: Salpida
Dec 1, 2017 · Salps are pelagic tunicates, which are widely distributed in oceans. They have holoplanktonic lifestyle and same as every holoplankton they are ...Missing: anatomy | Show results with:anatomy
[29]
molecular phylogeny of the Thaliacea | Journal of Plankton Research
In existing morphology-based classifications, the Order Salpida contains one family, the Salpidae, which comprises two subfamilies, the Cyclosalpinae and ...
[30]
Morphological and molecular characterization of salps (Thalia spp ...
Feb 28, 2014 · Novel cox1 barcoding primers developed in this study proved successful for the identification of Thalia salp species and generated much needed ...
[31]
A phylogenomic framework and timescale for comparative studies of ...
Apr 13, 2018 · Our study represents the most comprehensive phylogenomic dataset for the main tunicate lineages. It offers a reference phylogenetic framework and first ...
[32]
Phylogenetic analysis of phenotypic characters of Tunicata supports ...
Nov 13, 2019 · The endostyle is a conspicuous, trough along the ventral midline of the branchial basket. It contains glandular cells, ciliated cells and ...
[33]
Phylogeny of Tunicata inferred from molecular and morphological ...
The phylogeny of the Tunicata was reconstructed using molecular and morphological characters. Mitochondrial cytochrome oxidase I (cox1) and 18S rDNA ...
[34]
An updated 18S rRNA phylogeny of tunicates based on mixture and ...
Aug 5, 2009 · The high evolutionary rate of 18S rRNA sequences in Thaliacea also precluded firmly resolving the phylogenetic relationships among its three ...
[35]
A mid-Cambrian tunicate and the deep origin of the ascidiacean ...
Jul 6, 2023 · Additionally, tunicates have a poor fossil record, which includes only one taxon with preserved soft-tissues. Here we describe Megasiphon ...
[36]
Inferring Tunicate Relationships and the Evolution of the Tunicate ...
Mar 25, 2020 · Tunicate phylogenetic relationships remain poorly resolved across taxonomic levels. The approximately 3,000 species have historically been ...
[37]
First chromosome-level genome assembly of the colonial chordate ...
Sep 18, 2025 · Although Hox genes are colinear between cephalochordates and vertebrates, it is not the case for tunicates [70]. In the tunicate species studied ...
[38]
Salpa fusiformis | Zooplankton Guide
Cosmopolitan, with the widest distribution of all species of salp, occurs from 50° N and 45° S in the Pacific Ocean and from 70° N and 45° S in the Atlantic ...
[39]
[PDF] Distribution and Ecologic Aspects of Central Pacific Salpidae ...
It concerns the distribution and ecology of the salps collected during two cruises by the. "Hugh M. Smith" of the Pacific Oceanic. Fishery Investigations of the ...
[40]
Distribution, abundance, and reproductive stages of salps, doliolids ...
However, salps and doliolids show alternation of generations, which is a reproductive strategy that involves sexual reproduction in aggregates (blastozooids) ...
[41]
Reproductive biology, elemental composition and diel vertical ...
Jan 3, 2024 · Soestia is primarily an epipelagic salp species adapted to living in warm-temperate nutrient-depleted and more productive ocean regions.Materials And Methods · Results · Oceanographic Conditions
[42]
Periodic swarms of the salp Salpa aspera in the Slope Water off the ...
The Slope Water region south of New England in the western North Atlantic is one of these locations (Fig. 1). Sampling in this region during August 1975 ...
[43]
1: Spatial distribution of Salpa thompsoni and Ihlea racovitzai in the...
Ihlea racovitzai was more widely distributed, with maximum densities north of ~64 o S. S. thompsoni and I. racovitzai were recorded ...Missing: racovitzii | Show results with:racovitzii
[44]
Diel vertical migration bySalpa aspera and its potential for large ...
In mid-summer 1975 throughout the Western Slope Water of the North Atlantic Ocean, massive numbers ofSalpa aspera performed a diel vertical migration of at ...
[45]
Contrasting diel vertical migration patterns in Salpa fusiformis ...
Salps migrated to the surface during the day (reverse migration) at one station while they swam to surface layers at night (nocturnal migration) at the other ...
[46]
Size and stage specific patterns in Salpa thompsoni vertical migration
Salpa thompsoni were undergoing diel vertical migration between 450 and 100 m. Smaller individuals were the strongest migrators implying higher predation ...Missing: diurnal | Show results with:diurnal
[47]
(PDF) Diel vertical migration of the tunicate Salpa thompsoni in the ...
Aug 6, 2025 · This DVM pattern would have an advantage in increasing the feeding opportunities for salps. Although the longer stay at a bright depth may also ...
[48]
The non-selective Antarctic filter feeder Salpa thompsoni as ... - Nature
Jan 26, 2024 · The pelagic tunicate Salpa thompsoni is an important component of the Antarctic environment. Over the past few decades an expansion of this ...Missing: endemism | Show results with:endemism
[49]
Salps - Australian Antarctic Program
Aug 12, 2010 · Scientific names: Two species are common in Antarctic waters: Salpa thompsoni and Ihlea racovitzai. Physical description. Salps are gelatinous, ...Missing: endemism | Show results with:endemism
[50]
Salp blooms drive strong increases in passive carbon export in the ...
Feb 2, 2023 · Growth rates of blastozooids during SalpPOOP suggest a shorter life-cycle duration in the warmer waters (~10 °C) of the Chatham Rise, ranging ...
[51]
(PDF) Salp distribution and size composition in the Atlantic sector of ...
Aug 6, 2025 · Salp abundance and length frequency were measured during the large-scale CCAMLR 2000 Survey conducted in the Atlantic Sector of the Southern Ocean in the 1999/ ...
[52]
[PDF] Large salp bloom export from the upper ocean and benthic ...
Satellite estimates of chlorophyll and net primary production: 105. We used regionally optimized algorithms (Kahru et al. 2012) based on satellite. 106.
[53]
Salp metabolism: temperature and oxygen partial pressure effect on ...
Here I examine temperature and oxygen partial pressure effect on metabolism in blastozooids of Salpa fusiformis. Routine metabolic rates of 1.66 and 3.95 μmol ...
[54]
Hot Poop Temperature and Oxygen Partial Pressure Effects on Salp ...
Individual zooids were able to consume oxygen below detectable levels in respirometery chambers and recover when reoxygenated at both 17ºC and 10ºC. The ...
[55]
Grazing impact of salp (Tunicata, Thaliacea) assemblages in the ...
Salp assemblages ingested 0.01–3.5% of chlorophyll standing stock daily. The proportion of PP removed by salps was estimated to range from 0.1 to 24.5% day−1, ...
[56]
Salp blooms drive strong increases in passive carbon export in the ...
Feb 2, 2023 · The potential southward range expansion of salps may have important consequences for both the food web and the BCP, and biogeochemical models ...Missing: monitoring | Show results with:monitoring
[57]
Salp grazing: effects on phytoplankton abundance, vertical ...
Oct 5, 2025 · The major effects of salp grazing in the coastal environ- ment appear to be to deepen phytoplankton distributions and reduce biomass, rather ...
[58]
Small-scale effects of a river plume front on the distribution of salps ...
... salps and doliolids is probably due to their different feeding strategies. Salps have an almost neutral buoyancy, but they need to swim continuously for ...
[59]
Sensory ecology of salps (Tunicata, thaliacea): More questions than ...
Focussing on salps, this paper describes behavior that depends on sensory information, reviews the known structure and function of sensory receptors, and ...
[60]
https://academic.oup.com/plankt/article/37/6/1166/2380273
[61]
Prey taxonomy rather than size determines salp diets - ASLO - Wiley
Apr 22, 2019 · Dense swarms of salps have a high grazing impact that can deplete the photic zone of phytoplankton and export huge quantities of organic matter ...
[62]
clearance and ingestion rates of the salps Thalia democratica ...
The estimated clearance rate for all species based on disappearance of chlorophyll varied from 82 to 444 mL individual−1 day−1. Cell counts showed that T.Missing: liters | Show results with:liters
[63]
[PDF] High abundance of salps in the coastal Gulf of Alaska during 2011
Jun 10, 2016 · Large solitary individuals appeared capable of filtering 10–100 s of liters per day (Fig. 6). C. bakeri had lower filtration rates than S.
[64]
[PDF] Prey Size Spectra and Predator:Prey ratios of 7 Species of ... - bioRxiv
Feb 19, 2022 · Salps can have clearance rates >100,000 mL ind-1 day-1 (Madin ... by the average clearance rate for the size bins of particles expected to be ...
[65]
The Oceans Their Physics, Chemistry, and General Biology
Among the more or less obvious preying plankton feeders may be placed many fishes, notably herring, mackerel, sardines, and others of this type (p. ... salps ...<|separator|>
[66]
Gelatinous filter feeders increase ecosystem efficiency - PMC
Aug 23, 2024 · Food-web and isotope-based trophic analyses suggest that blooms of salps (gelatinous grazers) can increase ecosystem efficiency and energy transfer to large ...
[67]
Sinking of Gelatinous Zooplankton Biomass Increases Deep Carbon ...
Dec 17, 2019 · The short life span of most gelatinous zooplankton, from weeks up to 2 to 12 months (Ceh et al., 2015; Raskoff et al., 2003), suggests ...
[68]
New symbiotic associations of hyperiid amphipods (Peracarida) with ...
Hyperiid amphipods are holoplanktonic marine crustaceans that are known as temporary symbionts of different groups of gelatinous zooplankton.
[69]
Diverse microbial prey in the guts of gelatinous grazers revealed by ...
Mar 25, 2025 · Suspension feeding via mucous meshes or webs allows for the accumulation of detritus ... salp diets. Limnol Oceanogr 64(5):1996–2010. Article ...
[70]
The impact of salps (Salpa thompsoni) on the Antarctic krill ...
Oct 9, 2023 · All of the investigated krill population properties (abundance, mean length, and yearly egg production) were significantly impacted by the presence of salps.
[71]
Predictability of patches of neritic salps and doliolids (Tunicata ...
These intense phytoplankton blooms trigger rapid increases in asexual reproduction by salps and doliolids, resulting in mesoscale thaliacean patches containing ...
[72]
Distribution of life-history stages of the salp Thalia democratica in ...
Blooms of phytoplankton form along the continental shelf of eastern Australia as a result of the upwelling of cool nutrient-rich slope waters (Hallegraeff & ...
[73]
Salp blooms drive strong increases in passive carbon export in the ...
Particle export in salp waters is always higher, ranging 2- to 8- fold (average 5-fold), compared to non-salp locations, exporting up to 46% of primary ...
[74]
Our jelly-like relatives: Common misconceptions about salps
Sep 28, 2016 · But during a swarm, salps can cover an area up to 100,000 square km, with more than 5000 individuals per m3. Salps were also thought to be ...
[75]
Long-term changes in pelagic tunicates of the California Current
In the California Current, a swarm of Thalia democratica extended over 9000 km2 (Berner, 1967). Extensive regions of high salp abundances also have been ...Missing: historical | Show results with:historical
[76]
[PDF] Gelatinous zooplankton (ctenophores, salps and medusae)
May 3, 2017 · Salps were frequent prey on the slope and southern shelf. In contrast, medusae were consumed in coastal areas, slopes and the southern shelf.
[77]
What is a Salp? - Australian Museum
They are taxonomically closer to humans than jellyfish. Salps are classified in the Phylum chordata; they are related to all the animals with backbones.
[78]
Hydrodynamic advantages of swimming by salp chains - PMC - NIH
Aug 2, 2017 · We show that asynchronous swimming with multiple pulsed jets yields substantial hydrodynamic benefit due to the production of steady swimming velocities, which ...Missing: predator fragmentation rapid
[79]
Hurts My Brain – Salp Chain! | The Marine Detective
May 27, 2023 · When food is plentiful, they can quickly create more chains, and each salp can increase rapidly in size. ... Their lifespan depends on the ...
[80]
Salp blooms increase carbon export 5-fold in the Southern Ocean
Feb 8, 2022 · We show that blooms increased particle export by ~5-fold, and exported up to 46% of net primary production out of the euphotic zone.
[81]
Blooms of a key grazer in the Southern Ocean – An individual-based ...
Aug 6, 2025 · Hence, the risk of salp blooms in the WAP is likely to substantially increase. These findings highlight the importance for an improved ...
[82]
Towards an understanding of salp swarm dynamics - ICES Library
Mar 22, 2024 · A review and comparison of existing data is helping to define geographic hot spots for salp blooms as well as the necessary physical and biological precursors.
[83]
Salp contributions to vertical carbon flux in the Sargasso Sea
Salps are highly efficient filter feeders, with clearance rates up to several liters salp−1 ... Clearance rates of the salp Thalia democratica fed naturally ...Missing: per | Show results with:per
[84]
Krill and salp faecal pellets contribute equally to the carbon flux at ...
Dec 9, 2021 · ... feed on phytoplankton, based on a gut passage time of about 8 h. This pattern might decrease the efficiency of carbon export by krill FP.
[85]
[PDF] Distinguishing zooplankton fecal pellets as a component of the ...
The nitrogen isotope ratios of threonine and alanine statistically distinguished the zooplankton fecal pellets from literature-derived examples of phyto-.
[86]
Ammonia excretion rate of Salpa fusiformis Cuvier (Tunicata
The excretion rate increases allometrically with animal weight and exponentially with temperature.Missing: recycling | Show results with:recycling
[87]
dominance shift from krill to salps is associated with higher ...
Apr 3, 2020 · Hence, one might expect that an increasing occurance of salps at the WAP may lead to a shift towards higher N:P ratios in the dissolved nutrient ...
[88]
Antarctic pelagic ecosystems on a warming planet - ScienceDirect.com
Salp biomass in the Southern Ocean has increased and their distribution has expanded southward, intruding into areas historically dominated by Antarctic ...
[89]
Salp metabolism: temperature and oxygen partial pressure effect on ...
May 18, 2019 · There has been limited research on salp physiology and no studies that examine how changes in environmental factors such as temperature and ...
[90]
Effects of temperature on the swimming of salps (Tunicata, Thaliacea)
Jun 26, 1978 · This lag may be due to an actual physiological effect or to un- even warming. Since the salp was not swimming, the interior of the animal ...
[91]
Responses of Marine Organisms to Climate Change across Oceans
General expectations for biological and ecological responses to warming oceans include poleward distribution shifts, earlier spring events and delayed autumn ...
[92]
dominance shift from krill to salps is associated with higher ... - Nature
Apr 3, 2020 · Overall relation between dissolved N (µmol), dissolved P (µmol) as well as the N:P molar ratio and the salp and krill density (ind. 1000 m3) as ...
[93]
https://www.nature.com/articles/s41598-020-62829-8
[94]
New insight into Salpa thompsoni distribution via glider-borne ...
Jan 25, 2023 · There is a need to better understand the variability in salp densities and vertical distribution patterns in Antarctic waters to improve ...Missing: endemism | Show results with:endemism
[95]
Jelly Beans of the Sea - Meet the Marvelous Salp
Sep 29, 2022 · They are also capable of bioluminescence and are one of the brightest bioluminescent sea creatures out there. And as if this were not enough ...
[96]
https://www.thecricket.com/stories/jelly-beans-of-the-sea-meet-the-marvelous-salp%2C5153
[97]
First Voyage of Captain James Cook
First Voyage of Captain James Cook. (1768 - 1771). James Cook's first voyage circumnavigated the globe in the ship Endeavour, giving the botanists Joseph Banks ...
[98]
[PDF] Scripps Institution of Oceanography: Probing the Oceans, 1936 to ...
... William E. Ritter conducted his first summer field program, at Pacific Grove, California. 26 September 1903 - The Marine Biological Association of. San Diego ...
[99]
Acoustic backscattering from salp and target strength estimation
Very small plankton can be studied by using a high frequency echosounder when the number of individuals is big enough and when the body of the zooplankton ...Missing: imaging | Show results with:imaging
[100]
Rapid Evolutionary Rates and Unique Genomic Signatures ...
Abstract. A preliminary genome sequence has been assembled for the Southern Ocean salp, Salpa thompsoni (Urochordata, Thaliacea).
[101]
Latitudinal variation in plankton size spectra along the Atlantic ...
... salps could significantly contribute to the carbon export flux. Contrastingly, higher CR rates (.1000 mg C m22 day21) were found at the northern stations ...
[102]
As you may have heard, this summer Southern California had a salp ...
Sep 11, 2024 · On the summer CalCOFI cruise, we saw huge numbers and patches of high density of these cool gelatinous organisms nearshore, and we couldn't help ...
[103]
Massive salp outbreaks in the inner sea of Chiloé Island (Southern ...
Aug 6, 2025 · First outbreaks were recorded during February 2010 when Ihlea magalhanica reached up to 654,000 ind m -3 close to the net pens in Maillen Island ...
[104]
[PDF] bioprospecting for novel psychrophilic enzymes from the Antarctic ...
Initial results from 16S analysis of the gut microbiome indicate the presence of a Psychrobacter species which may aid in cellulytic digestion within salp ...
[105]
Patterns of suspended and salp‐ingested microplastic debris in the ...
Nov 27, 2019 · Mini-microplastics were found in nearshore waters and all salps ingested them. Every salp had ingested plastic, with blastozooids having higher ...
[106]
Predicting Harmful Algal Blooms Using Explainable Deep Learning ...
The Transformer model's architecture, leveraging self-attention mechanisms and positional encodings, can be a powerful tool for forecasting harmful algal blooms ...